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Phylum XVI. Tenericutes Murray 1984A, 356VP (Effective Publication: Murray 1984B, 33.) Da N I E L R

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Phylum XVI. Tenericutes Murray 1984A, 356VP (Effective Publication: Murray 1984B, 33.) Da N I E L R Phylum XVI. Tenericutes Murray 1984a, 356VP (Effective publication: Murray 1984b, 33.) DANIEL R. BR OWN Ten.er¢i.cutes. L. adj. tener tender; L. fem. n. cutis skin; N.L. fem. n. Tenericutes prokaryotes of a soft pliable nature indicative of a lack of a rigid cell wall. Members of the Tenericutes are wall-less bacteria that do not syn- the organisms are evolutionarily related to certain clostridia, thesize precursors of peptidoglycan. the absence of a cell wall cannot be equated with Gram reac- tion positivity or with other members of the Firmicutes. It is Further descriptive information unfortunate that workers involved in determinative bacteriology The nomenclatural type by monotypy (Murray, 1984a) is the have a reference in which wall-free prokaryotes are described as class Mollicutes, which consists of very small prokaryotes that Gram-positive bacteria” (Tully, 1993a). Despite numerous valid are devoid of cell walls. Electron microscopic evidence for the assignments of novel species of mollicutes to the Tenericutes dur- absence of a cell wall was mandatory for describing novel species ing the intervening years, the class Mollicutes was still included of mollicutes until very recently. Genes encoding the pathways in the phylum Firmicutes in the most recent revision of the Taxo- for peptidoglycan biosynthesis are absent from the genomes of nomic Outline of Bacteria and Archaea (TOBA), which is based more than 15 species that have been annotated to date. Some solely on the phylogeny of 16S rRNA genes (Garrity et al., 2007). species do possess an extracellular glycocalyx. The absence of The taxon Tenericutes is not recognized in the TOBA, although a cell wall confers such mechanical plasticity that most molli- paradoxically it is the phylum consisting of the Mollicutes in the cutes are readily filterable through 450 nm pores and many spe- most current release of the Ribosomal Database Project (Cole cies have some cells in their populations that are able to pass et al., 2009). Mollicutes are specifically excluded from the most through 220 nm or even 100 nm filters. However, they may vary recently emended description of the Firmicutes in Bergey’s Manual in shape from coccoid to flask-shaped cells or helical filaments of Systematic Bacteriology (2nd edition, volume 3; De Vos et al., that reflect flexible cytoskeletal elements. 2009) on the grounds of their lack of rigid cell walls plus analyses of strongly supported alternative universal phylogenetic markers, Taxonomic comments including RNA polymerase subunit B, the chaperonin GroEL, To provide greater definition and formal nomenclature for several different aminoacyl tRNA synthetases, and subunits of vernacular names used in the 8th edition of Bergey’s Manual of F0F1-ATPase (Ludwig et al., 2009; Ludwig and Schleifer, 2005). Determinative Bacteriology (Bergey VIII; Buchanan and Gibbons, The taxonomic dignity of Tenericutes bestowed by its original 1974), Gibbons and Murray (1978) proposed that the higher formal validation, and upheld by a quarter of a century of valid taxa of prokaryotes be subdivided primarily according to the descriptions of novel species of mollicutes, has therefore been presence and character, or absence, of a rigid or semirigid cell respected in this volume of Bergey’s Manual. wall as reflected in the determinative Gram reaction. Similar Type order: Mycoplasmatales Freundt 1955, 71AL emend. Tully, to the non-hierarchical groupings of Bergey VIII, which were Bové, Laigret and Whitcomb 1993b, 382. based on a few readily determined criteria, the “wall-deficient” organisms grouped together in the first edition of The Prokary- References otes included the mollicutes (Starr et al., 1981). While acknowl- Buchanan, R.E. and N.E. Gibbons (editors). 1974. Bergey’s Manual of edging the emerging 16S rRNA-based evidence that indicated Determinative Bacteriology, 8th edn. Williams & Wilkins, Baltimore. a phylogenetic relationship between mollicutes and certain Cole, J.R., Q. Wang, E. Cardenas, J. Fish, B. Chai, R.J. Farris, A.S. Kulam- Gram-stain-positive bacteria in the division Firmicutes, Murray Syed-Mohideen, D.M. McGarrell, T. Marsh, G.M. Garrity and J.M. (1984b) proposed the separate division Tenericutes for the stable Tiedje. 2009. The Ribosomal Database Project: improved alignments and distinctive group of wall-less species that are not simply an and new tools for rRNA analysis. Nucleic Acids Res. 37: (Database obvious subset of the Firmicutes. issue): D141–D145. The approved divisional rank of Tenericutes and the assignment De Vos, P., G. Garrity, D. Jones, N.R. Krieg, W. Ludwig, F.A. Rainey, K.H. of class Mollicutes as its nomenclatural type (Murray, 1984a) were Schleifer and W.B. Whitman. 2009. In Bergey’s Manual of Systematic adopted by the International Committee on Systematic Bacte- Bacteriology, 2nd edn, vol. 3. Springer, New York. Freundt, E.A. 1955. The classification of the pleuropneumoniae group riology’s Subcommittee on the Taxonomy of Mollicutes (Tully, of organisms (Borrelomycetales). Int. Bull. Bacteriol. Nomencl. Taxon. 1988) and subsequent valid taxonomic descriptions assigned 5: 67–78. novel species of mollicutes to the Tenericutes. However, the second Garrity, G.M., T.G. Lilburn, J.R. Cole, S.H. Harrison, J. Euzéby and B.J. (1992) and third (2007) editions of The Prokaryotes described the Tindall. 2007. The Taxonomic Outline of the Bacteria and Archaea, mollicutes instead as Firmicutes with low G+C DNA. The Subcom- Release 7.7, Part 11 – The Bacteria: Phyla Planctomycetes, Chlamyd- mittee considered this to be an unfortunate grouping: “While iae, Spirochaetes, Fibrobacteres, Acidobacteria, Bacteroidetes, Fusobacteria, 567 568 PHYLUM XVI. TENERICUTES Verrucomicrobia, Dictyoglomi, Gemmatomonadetes, and Lentisphaerae. pp. Starr, M.P., H. Stolp, H.G. Trüper, A. Balows and H.G. Schlegel (editors). 540–595. (http://www.taxonomicoutline.org/). 1981. The Prokaryotes. Springer, Berlin. Gibbons, N.E. and R.G.E. Murray. 1978. Proposals concerning the Tully, J.G. 1988. International Committee on Systematic Bacteriol- higher taxa of bacteria. Int. J. Syst. Bacteriol. 28: 1–6. ogy, Subcommittee on the Taxonomy of Mollicutes, Minutes of the Ludwig, W. and K.H. Schleifer. 2005. Molecular phylogeny of bacte- Interim Meeting, 25 and 28 August 1986, Birmingham, Alabama. Int. ria based on comparative sequence analysis of conserved genes. In J. Syst. Bacteriol. 38: 226–230. Microbial Phylogeny and Evolution, Concepts and Controversies, Tully, J.G. 1993a. International Committee on Systematic Bacteriol- (edited by Sapp). Oxford University Press, New York, pp. 70–98. ogy, Subcommittee on the Taxonomy of Mollicutes, Minutes of the Ludwig, W., K.H. Schleifer and W.B. Whitman. 2009. Revised road map to Interim Meetings, 1 and 2 August, 1992, Ames, Iowa. Int. J. Syst. the phylum Firmicutes. In Bergey’s Manual of Systematic Bacteriology, 2nd Bacteriol. 43: 394–397. edn, vol. 3, The Firmicutes (edited by De Vos, Garrity, Jones, Krieg, Lud- Tully, J.G., J.M. Bové, F. Laigret and R.F. Whitcomb. 1993b. Revised wig, Rainey, Schleifer and Whitman). Springer, New York, pp. 1–13. taxonomy of the class Mollicutes – proposed elevation of a monophyl- Murray, R.G.E. 1984a. In Validation of the publication of new names etic cluster of arthropod-associated mollicutes to ordinal rank (Ento- and new combinations previously effectively published outside the moplasmatales ord. nov.), with provision for familial rank to separate IJSB. List no. 15. Int. J. Syst. Bacteriol. 34: 355–357. species with nonhelical morphology (Entomoplasmataceae fam. nov.) Murray, R.G.E. 1984b. The higher taxa, or, a place for everything…? In from helical species (Spiroplasmataceae), and emended descriptions Bergey’s Manual of Systematic Bacteriology, vol. 1 (edited by Krieg of the order Mycoplasmatales, family Mycoplasmataceae. Int. J. Syst. and Holt). Williams & Wilkins, Baltimore, pp. 31–34. Bacteriol. 43: 378–385. Class I. Mollicutes Edward and Freundt 1967, 267AL DANIEL R. BR OWN , MEGHAN MAY , JANET M. BR A db U R Y AN D KA R L -ER I K JOHANSSON Mol¢li.cutes or Mol.li.cu¢tes. L. adj. mollis soft, pliable; L. fem. n. cutis skin; N.L. fem. pl. n. Mollicutes class with pliable cell boundary. Very small prokaryotes totally devoid of cell walls. Bounded pathogens of humans, animals, insects, or plants. Genome sizes by a plasma membrane only. Incapable of synthesis of pepti- range from 580 to 2200 kbp, among the smallest recorded in doglycan or its precursors. Consequently resistant to penicillin prokaryotes. The genomes of more than 20 species have been and its derivatives and sensitive to lysis by osmotic shock, deter- completely sequenced and annotated to date (Table 135). The gents, alcohols, and specific antibody plus complement. Gram- G+C content of the DNA is usually low, ~23–34 mol%, but in some stain-negative due to lack of cell wall, but constitute a distinct species is as high as ~40 mol% (Bd, Tm). Can be distinguished from phylogenetic lineage within the Gram-stain-positive bacteria other bacteria in having only one or two rRNA operons (one spe- (Woese et al., 1980). Pleomorphic, varying from spherical or cies of Mesoplasma has three) and an RNA polymerase that is resis- flask-shaped structures to branched or helical filaments. The tant to rifampin. The 5S rRNA contains fewer nucleotides than coccoid and flask-shaped cells usually range from 200–500 nm that of other bacteria and there are fewer tRNA genes. In some in diameter, although cells as large as 2000 nm have been seen. genera, instead of a stop, the UGA codon encodes tryptophan. Replicate by binary fission, but genome replication may pre- Plasmids and viruses (phage) occur in some species. cede cytoplasmic division, leading to the formation of multi- Type order: Mycoplasmatales Freundt 1955, 71AL emend. Tully, nucleated filaments. Colonies on solid media are very small, Bové, Laigret and Whitcomb 1993, 382. usually much less than 1 mm in diameter. The organisms tend to penetrate and grow inside the solid medium.
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